the pollen flora of faiyum, egypt i- archichlamydeae · 2020. 9. 3. · 2 eman m. shamso and fujiki...

3 Taeckholmia 32 (2012): 1-40

The Pollen Flora of Faiyum, Egypt I- Archichlamydeae

Eman M. Shamso¹ and Fujiki Toshiyuki²

¹ The Herbarium, Botany Department, Faculty of Science, Cairo University, Giza 12613 Egypt; e-mail: [email protected] ²International Research Center for Japanese Studies, Japan; e-mail: [email protected]

Eman M. Shamso and Fujiki Toshiyuki. 2012. The Pollen Flora of Faiyum, Egypt I- Archichlamydeae. Taeckholmia 32: 00-00.

The aim of this investigation is dealing with determination of pollen micro- and macro-morphological characters of Egyptian taxa of subclass Archichlamydeae to provide palynological information and an identification key adapted for use with archeobotanical materials. The pollen of 41 Egyptian taxa belongs to 18 families representing the current vegetation of Faiyum area described and illustrated with both LM and SEM micrographs. On the basis of pollen units, number and type of apertures, exine patterns were great diagnostic significance among the examined taxa. Seven major pollen types and 18 subtypes included in two groups (complex and single pollen groups) were recognized. Pollen types: Polyads, triporate, tetra- hexaporate and pantobrevicolpate are characterized to Mimosoideae, Urticaceae, Fumariaceae and Portulacaceae respectively; while pantoporate, tricolpate and tricolporate pollen types show highly diversity in pollen characters, comprising 16 pollen subtypes distributed among 14 families. A description, illustriation in the form of scanning electron micrographs and a key for identification of the different taxa is provided.

Received 1 November 2012, Accepted 16 November 2012

mailto:[email protected]

mailto:[email protected]

Eman M. Shamso and Fujiki Toshiyuki 2

Key words: Archichlamydeae, Faiyum, Identification key, Pollen Flora, Pollen types, SEM micrographs.

Introduction

Western Desert of Egypt is one of the main geomorphologic units, which extends from the Mediterranean Coast in the north to the Egyptian- Sudanese border in the south (ca. 1073km) and from the Nile Valley in the east to the Egyptian-Libyan border in the west (width ranges between 600- 850 km), i.e. it covers about > of Egypt (El Hadidi 2000).

Faiyum province is one of the depressions of the Western Desert of Egypt. Being the nearest to the Nile Valley and after being connected with the River Nile by a large irrigation canal (Bahr Yusuf), The Faiyum Depression is considered as apart of the Nile Region (Nile Faiyum). The depression has a total area of about 1700 km² (Zahran & Willis 2009).

The plants of the Nile system of Egypt (about 553 species) represent 29.9% of the total flora of Egypt. About 126 species are not recorded elsewhere in Egypt; of these species 64 characterize the Nile Faiyum (Zahran & Willis 1992).

Täckholm (1974) recorded about 129 taxa representing subclass Archichlamydeae restricted to the Nile Valley area. The current vegetation of the Faiyum area was the subject of several recent accounts, Abd El Ghani (1985) recorded 158 species in the northern district of Faiyum area, which constitute the major bulk of the weed assemblages associated with the crops of the farm land.

El Fayoumi (1996) studied the main features of the current vegetation in El Gharag El Sultani area which constitute the southern districts of the farm land in the Faiyum depression, he recorded 121 species, among which 115 species are weeds of the farm land.

Saad & Sami 1967, El shenbary 1985, Ritchie 1985 and Ayyad et al. 1992b studied the fossil pollen content in different areas of Egypt. Mehringer et al. 1979 carried out a pollen analytical study in the lake Qarun area and elucidated the history of Birket Qarun and Fayium over the last 325 years, while El Hadidi & El Fayoumi 1997 recognized 86 pollen types grouped under 21 pollen classes in El Gharag El Sultani area using light microscopy (LM), he provided artificial key and brief descriptions for the pollen types.

The present investigation is dealing with determination of pollen micro- and macro-morphological characters of 41 taxa of subclass Archichlamy-

3 The Pollen Flora of Faiyum, Egypt I- Archichlamydeae

deae (sensu Engler 1964) representing the current vegetation of Faiyum area, using light and scanning electron microscopy. This investigation aims to provide palynological information of all examined taxa, to cluster the taxa involved according to resemblance in pollen characters and to present an identification key based on pollen morphology adapted for use with archeobotanical material.

Materials and Methods

Pollen samples of 41 species representing 37 genera and 18 families had been collected from herbarium specimens kept in the Cairo University Herbarium (CAI) and fresh materials collected from the field (Table 1). All materials were taken from mature anthers of flowering buds. List of specimens investigated is in accordance with the system of Engler 1964; nomenclature of the taxa was updated according to Boulos (2009). Voucher specimens and pollen slides were kept in Cairo University Herbarium (CAI).

Pollen grains for light microscope (LM) and scanning electron microscope (SEM) were prepared by 10% KOH treatment and Erdtman's acetolysis method (Erdtman 1960). For light microscope, samples were dehydrated with ethanol series (60, 80 & 99.5%) after acetolysis method, then substituted with xylene; and were mounted in eukitt on the slide. Light microscopic observations were made by using Nikon Eclipse E 600, the pollen sizes were measured at 600 x magnification, and photographs of the samples taken at 400 x. As for scanning electron microscopic observations, samples were fixed and conductive coated with 2% Osmium tetroxide solution after acetolysis, then dehydrated with ethanol series (60, 80, and 99.5%) and substituted with xylene. The acetolyzed pollens were spattered with gold – palladium for about five minutes on the sample stand. Scanning electron microscope observations were made using JEOL JSM-6300F (the laboratory of International Research Center, Japan). The SEM images were taken at acceleration voltage of 3-4kv. Comparative analysis of pollen morphology and masurements of qualitative parameters of pollen grains as well as pollen preparations were done at the laboratory of International Research Center for Japanese Studies (Japan) and the Herbarium of Cairo University (Egypt).

All measurements are based on at least 15-20 pollen grains for each specimen, various pollen characters viz: shape, size, aperture type and exine


sculpture were determined. The terminology used follows Erdtman (1952), Praglowski & Punt (1973), Praglowski & Raj (1979) and Punt et al. (2007).

Results

The palynological data of the investigated taxa are summarized in Table (2). The examination of the available pollen materials revealed the presence of two groups based on pollen units:

A. Complex grains group: Pollen grains released in polyads, composed of 16 or 32 grains in regular arrangement. This group represented by a unique major pollen type known as Polyads type, which subdivided into two subtypes.

B. Single grains group: pollen grains released in monads, usually spheroidal or varied from prolate-spheroidal to prolate or suboblat. Six major pollen types can be distinguished based on the shape and number of apertures viz. triporate, tetra-hexa porate, pantoporate, panto-brevicolpate, tricolpate and tricolporate types. Among these major types, 16 pollen subtypes were recognized based on the exine sculpture, pollen shape and size and other pollen characters.

TABLE (1). Collection data of the investigated taxa arranged in accordance with the

system of Engler 1964 (arranged alphabetical). Family Examined taxa Locality

Aizoaceae Mesembryanthemum crystallinum L.

Burg el Arab, Mariut; 22/3/1956; I. El Sayed s.n. (CAI).

Amaranthaceae Alternanthera sessilis (L.) DC. Wadi El Rayan; 15/4/2000; Boulos s.n.(CAI). Amaranthus graecizans L. Tamiya District, Kafr Mahfous; 12/11/1982; Abd El

Ghani 4423(CAI).

Capparaceae Capparies decidus (Forssk.) Edgew.

Wadi El Allagi (upstream part); 25/1/1963; Täckholm et al. s.n.(CAI).

Caryophyllaceae Silene rubella L. Near Qarun lake; 29/2/1988; E. Shamso s.n.(CAI). Spergularia marina (L.)

Bessler Faiyum; 29/4/1979; A. Hosny s.n.(CAI)

Vaccaria hispanica (Mill.) Rauschert

Faiyum; 12/1/1952; El Hadidi s.n.(CAI).

Chenopodiaceae Arthrocnemum macrostachym (Moric.) K. Koch.

In the salt marshes near Lake Quroun, Faiyum; 6/12/1965; Botany Depart. Excursion s.n.(CAI).

Beta vulgaris L. Kom Aushim, Faiyum; 18/3/1977; Täckholm et al. s.n.(CAI).

Chenopodium album L. Faiyum; 12/1/1959; A. Amin s.n.(CAI). Suaeda aegyptiaca (Hasselq.)

Zohary 8 km from Faiyum on the Lahun-Faiyum road; 6/10/1967; Täckholm et al. s.n.(CAI).

Cruciferae Brassica nigra (L.) Koch Faiyum; Kom Aushim; 4/3/1962; Täckholm et al. s.n.(CAI).

Sinapis arvensis L. Faiyum; Kom Aushim; Täckholm et al. s.n.(CAI). Fumariaceae Fumaria bracteosa Pomel Ikingi Marut; 3/4/1977; Kosinova et al. s.n.(CAI).


Leguminoseae Acacia nilotica (L.) Delil. Nag El –Ismailia, Ballana; 22/12/1963; Boulos s.n. (CAI).

Alhagi graecorum Boiss. Beheira province, Abu Hummus; 24/11/1987; Amer s.n.(CAI).

Faidherbia albida (Delil.) A. Chev.

River bank, Khartum, Sudan; 9/12/1954; Unknown (CAI).

Lathyrus hirsutus L. Sennuris, Faiyum; 5/5/1967; El Hadidi s.n.(CAI). Lotus halophilus Boiss. &

Spruner Beheira province, Idku; 29/4/1987; Amer s.n.(CAI).

Lotus pusillus Viv. 25 km W of Syrte; 8/3/1968; Boulos s.n.(CAI). Medicago intertexta (L.) Mill. Faiyum; 11/1/1972; I El Sayed s.n.(CAI). Melilotus indicus Behiera Province, Kom Hamada; 17/1/1987; Amer

s.n.(CAI). Sesbania sesban (L.) Merr. Khour Rl Mlagi; 21/3/1962; Abdalla et al. s.n.(CAI). Trifolium resupinatum L. Baharia Oasis, El Heig Aui Heg; 13/9/1971; M. Iwan

s.n.(CAI). Vicia monantha Retz. Kharga & Dakhla Oasis, Dakhla: Mut; 13/2/1952;

Täckholm & Kassas s.n.(CAI). Malvaceae Malva parviflora L. Faiyum; 4-1-1991; Araffa s.n.(CAI).

Oxalidaceae Oxalis corniculata L. Bawiti, Ain Miftilla, in a date palm grove; 17/7/1978; Abd El Ghani s.n.(CAI).

Polygonaceae Emex spinosa (L.) Campd El Faiyum; 31/1/1975; Hossny et.al s.n.(CAI). Persicaria salicifolia (Brouss.

Ex Willd.) Assenov Samouh cultivations behind Nuzha Gardens, Alexandria; 23/3/1956; Täckholm & El Hadidi s.n.(CAI).

Polygonum equsetiforme Sm. Qouran lake, El Faiyum; 24/4/1972; Zahran s.n.(CAI). Rumex dentatus L. Ali Bey SW of Madinet El Faiyum; 21/11/1926; G.

Täckholm s.n.(CAI). Rumex pectus Forssk. Cairo-Alexandria desert road; 8/3/1978; A. Soliman

s.n.(CAI). Portulacaceae Portulaca oleracea L. Near Qarun lake; 29/2/1988; E. Shamso s.n.(CAI). Salicaceae Salixmucronata Thunb. Zohria Garden; 15/1/1959; Hefnawy s.n.(CAI).

Salvadoraceae Salvadora persica L. Wadi El Balie, Hurghda district; 5/9/1960; Täckholm et al.s.n.(CAI).

Tamaricaceae Tamarix nilotica (Ehrenb.) Bunge

Kom Aushim, Faiyum; 21/9/1959; Täckholm s.n.(CAI).

Tamarix tetragyna Ehrenb. Kom Aushim, Faiyum; 11/3/1987; E. Shamso s.n.(CAI).

Thymelaeaceae Thymelaea hirsute (L.) Endl. Mariut, N of Amria station; 25/3/1927; G. Täckholm s.n.(CAI).

Urticaceae Urtica urens L. Beheira Province, Damanhour; 6/3/1988; Amer s.n.(CAI). Zygophyllaceae Fagonia Arabica L. Faiyum; 11/3/1987; E. Shamso s.n.(CAI).

Zygophyllum coccineum L. El Faiyum; 11/5/1973; Sisi et al. (CAI).

Description of Pollen types A. Complex grains group 1. Polyads type:

Polyads biconvex and circular to slightly elliptic in outline, medium to large, mean polar axis ranges from 24.8-47.3Pm and mean equatorial diameter ranges from 39.4-101.9 Pm; composed of 16 or 32 grains in a regular arrangement. The pollen grains are heteropolar, pyramidal or polygonal, colporate or indistinct aperture, exine sculpture foveolate or subrugulate. This type can be divided into two subtypes as follow:

6 Eman M. Shamso and Fujiki Toshiyuki

TABLE (2). The pollen morphological characters of the examined taxa of subclass Archichlamydeae ( values in parentheses represent average lengths).

Family

Taxa Polar axis P (-m)

Equatoria l axis E

(-m)

P/E ratio

Shape in Equatorial

view

Shape in Polar view

Aperture type

Colpus shape

Ora shape Exine sculpture

Pattern

Aizoaceae Mesembryanthemum crystallinum

19.2 – 21.7 (21.1±0.8)

18.4 – 21.7 (19.5±0.9)

0.96-1.18 (1.08±0.06)

Spheroidal or subprolate Circular Tricolporate Narrowly

oblong Sunken & Lolongate

Micro-perforate- granulate

Amaranthaceae Alternanthera sessilis

13.4-16.7 (14.6±1.0)

13.4-16.7 (14.6±1.0) 1.00 Spheroidal Octagonl Pantoporate --- Flat & Circular Metareticulate

(Fenestrate)

Amaranthus graecizans

18.4-23.4 (20.0±1.0)

18.4-23.4 (20.0±1.0)

1.00

Spheroidal

Circular

Pantoporate

--- Sunken, Circular

& operculate

Granulate to granulate with faint perforate

Capparaceae

Capparies decidus 15.0-20.9

(18.8±1.4) 13.4-17.5 (15.8±1.1)

0.90-1.39 (1.19±0.90)

Spheroidal subprolate or

prolate

Circular

Tricolporate

Fusiform

Flat & Lolongate

Perforate

Caryophyllaceae Silene rubella 31.7-34.5 (34.5±0.2)

31.7-34.5 (34.5±0.2) 1.00 Spheroidal Circular Pantoporate --- ±Sunken, Circular

& operculate Microechinate- anulopunctate

Spergularia marina 15.9-21.7

(18.7±1.4) 17.5-23.4 (19.9±1.6)

0.85-1.09 (0.94±0.06)

Suboblate or

Spheroidal

Circular

Tricolpate

Oblong

--- Microechinate-

perforate

Vaccaria hispanica 35.1-40.9 (38.3±1.3)

35.1-40.9 (38.3±1.3) 1.00 Spheroidal Circular Pantoporate --- ±Sunken, Circular

& operculate Microechinate- anulopunctate

Chenopodiaceae Arthrocnemum

macrostachym 20.0-23.4 (21.0±0.9)

20.0-23.4 (21.0±0.9)

1.00

Spheroidal

Circular

Pantoporate


& operculate

Granulate – psilate

Beta vulgaris 16.7-20.9 (18.6±1.1)

16.7-20.9 (18.6±1.1) 1.00 Spheroidal Circular Pantoporate --- Sunken, Circular

& operculate Granulate – psilate

Chenopodium album 21.7-30.1 (23.8±2.0)

21.7-30.1 (23.8±2.0) 1.00 Spheroidal Circular Pantoporate --- Sunken, Circular

& operculate Granulate – psilate

Suaeda aegyptiaca 21.7-26.7

(24.5±1.3) 21.7-26.7 (24.5±1.3)

1.00

Spheroidal

Circular

Pantoporate


& operculate

Granulate – psilate

Cruciferae

Brassica nigra

21.7-25.1 (23.6±1.4)

21.7-26.1 (23.7±1.2)

0.93-1.2 (1.00±0.06)

Spheroidal

Obtusely triangular

tricolpate

Fusiform

---

Reticulate with free standing columellae in

lumina


Sinapis arvensis

21.7-26.7 (23.9±1.1)

23.4-28.4 (26.5±1.2)

0.82-1.00 (0.9±0.04)

Oblate- sheroidal or suboblate

Obtusely triangular

Tricolpate

Fusiform

---

Reticulate with free standing columellae in

lumina

Fumariaceae Fumaria bracteosa 21.7 - 26.7 (24.9±1.3)

21.7 - 26.7 (24.9±1.3) 1.00 Sheroidal Circular Tetra- to

hexa-porate --- Raised, circular & aspidate

Verrucate - perforate

Leguminosae Acacia nilotica 23.4-26.7 (24.8±0.7)

36.7-43 (39.4±1.7)

0.58-0.68 (0.63±0.03)

Oblate Circular Colporate Y-shaped indistinct foveolate

Alhagi graecorum

15.0-20.0 (16.5±1.3)

13.4-15.9 (14.1±0.8)

1.00-1.50

(1.18±0.10)

Prolate- spheroidal,

subprolate or prolate

Sub-

circular

Tricolporate

Slightly fusiform

Raised & lolongate

Microreticulate

Faidherbia albida 4.18-51.8

(47.3±4.1)

91.9-115.2 (101.9±6.4

)

0.42-0.53 (0.47±0.04)

Prolate or oblate

Circular or ellipsoidal

indistincate

---

---

Subrugulate

Lathyrus hirsutus 31.7-41.8 (37.6±2.1)

22.5-30.1 (26.3±1.4)

1.22-1.64 (1.51±0.11)

Subprolate or Prolate

Obtusely triangular Tricolporate Narrowly

oblong Flat & lolongate Faint rugulate

Lotus halophilus 15.0-18.4 (16.8±0.8)

10.0-12.5 (11.6±0.6)

1.33-1.57 (1.44±0.06) Prolate Circular Tricolporate Slit-like Flat & lolongate Rugulate-fossulate

Lotus pusillus 14.2-18.4 (16.1±1.0)

10.0-12.5 (11.1±0.7)

1.29-1.58 (1.44±0.07) Prolate Circular Tricolporate Slit-like Flat & lolongate Rugulate-fossulate

Medicago intertexta 26.7-32.6 (28.8±1.4)

25.1-31.7 (27.9±1.6)

0.94-1.13 (1.03±0.05) Spheroidal Sub-

circular Tricolporate Narrowly oblong

Flat & lolongate Rugulate-fossulate

Melilotus indicus 20.9-26.7 (24.2±1.3)

16.7-21.7 (18.4±1.4)

1.17-1.50 (1.32±0.08)


Sub- circular Tricolporate Slightly

fusiform Raised &lolongate Microreticulate

Sesbania sesban 23.4-26.7 (24.2±1.0)

21.7-26.7 (24.7±1.4)

0.91-1.14 (0.98±0.06) Spheroidal Sub-

circular Tricolporate fusiform Raised &lolongate Perforate

Trifolium resupinatum

25.1-28.4 (26.3±0.9)

19.2-23.4 (20.6±1.0)

1.15-1.39 (1.28±0.06)


Sub- circular Tricolporate Narrowly

oblong ±Raised & lolongate

Reticulate- fossulate

Vicia monantha 31.7-36.7

(34.1±1.4) 20.0-23.4 (22.1±0.9)

1.36-1.79 (1.54±0.11)

Prolate or perprolate Sub-

circular

Tricolporate Narrowly

oblong ±Raised & lolongate

Reticulate- rugulate

Malvaceae Malva parviflora 75.2-85.2 (80.3±3.0)

75.2-85.2 (80.3±3.0) 1.00 Spheroidal Circular Pantoporate --- Flat & circular Sharp echinate &

perforate

Oxalidaceae Oxalis corniculata 23.4-36.7 (31.1±2.5)

26.7-36.7 (32.7±2.1)

0.88-1.05 (0.95±0.05) Spheroidal Circular Tricolpate fusiform --- Microreticulate to

perforate

Polygonaceae

Emex spinosa 21.7-26.7 (24.6±1.2)

25.1-28.4 (26.4±1.2)

0.87-1.10 (0.94±0.04)

Suboblate or spheroidal

Circular

Tricolporate Very short

& fusiform Sunken & lolongate

Fossulate- perforate with

granules

8 Eman M. Shamso and Fujiki Toshiyuki

Persicaria salicifolia 36.7-51.8 (45.8±3.8)

36.7-51.8 (45.8±3.8) 1.00 Spheroidal Circular Pantoporate --- Sunken & circular Lopho-reticulate

Polygonum equsetiforme

28.4-33.4 (30.0±1.2)

21.7-25.9 (23.2±0.9)

1.20-1.43 (1.31±0.05)


Sub- circular Tricolporate oblong Sunken &

lolongate Microperforate with granules

Rumex dentatus 21.7-26.7

(24.1±1.3) 25.1-28.4 (26.2±1.3)

0.87-1.07 (0.93±0.05)

Suboblate or oblate-

spheroidal

Circular Tricolporate

Slit-like Sunken & lolongate


granules

Rumex pectus 18.4-21.7

(20.4±0.8) 20.0-23.4 (21.3±0.9)

0.86-1.00 (0.96±0.04)


spheroidal

Sub- circular

Tricolporate Slit-like Sunken &

lolongate


granules

Portulacaceae Portulaca oleracea 48.2-58.5 (53.6±3.2)

48.2-58.5 (53.6±3.2) 1.00 Spheroidal Circular Pantocolpate Slightly

fusiform --- Microechinate - anulopunctate

Salicaceae

Salix mucronata 16.7-23.4

(20.8±1.6) 15.0-20.0 (17.1±1.4)

1.09-1.47 (1.22±0.12)

Spheroidal, or prolate

Obtusely triangular

Tricolporate

Fusiform

Sunken & circular

Uneven reticulate with columellae in

lumina

Salvadoraceae Salvadora persica 10.9-15.6 (12.9±1.3)

10.9-10.0 (12.6±1.2)

0.88-1.21 (1.02±0.08)

Spheroidal or subprolate

Obtusely triangular Tricolporate Widely

oblong Sunken & lalongate perforate

Tamaricaceae Tamarix nilotica 13.4-16.4 (14.2±0.9)

13.4-16.7 (14.6±0.9)

0.89-1.15 (0.97±0.08)


Obtusely triangular Tricolpate fusiform --- microreticulate

Tamarix tetragyna 15.0-18.4 (16.9±0.9)

15.0-19.2 (16.8±1.1)

0.91-1.22 (1.00±0.08)


Obtusely triangular Tricolpate fusiform --- microreticulate

Thymelaeaceae Thymelaea hirsute 21.7-25.1 (22.7±1.1)

21.7-25.1 (22.7±1.1) 1.00 Spheroidal Circular Pantoporate --- Sunken & circular Croton pattern

Urticaceae

Urtica urine 11.7-14.2

(13.4±1.1) 11.7-16.7 (14.7±1.3)

0.84-1.00 (0.91±0.05)


spheroidal

Circular

Triporate

--- Sunken, annulate

& operculate Densely spaced microechinate

Zygophyllaceae Fagonia arabica 15.0-20.9 (17.8±1.6)

13.4-18.4 (16.3±1.3)

0.95-1.33 (1.10±0.08)

Spheroidal or subprolate Circular Tricolporate Fusiform Flat & lalongate Microreticulate

Zygophyllum coccineum

9.2-10.0 (9.9±0.3)

9.0-10.2 (9.9±0.9)

0.92-1.20 (1.02±0.6)

Spheroidal or subprolate Circular Tricolporate Fusiform Flat & lalongate Microreticulate


1a. Acacia subtype Polyads circular, medium sized (23.4-26.7 Pm x 36.7-43.4 Pm),

composed of 16 grains. Eight grains in the centre, placed in two planes, each plane four grains surrounded by eight peripheral grains. Pollen grains colporate, colpi Y-shaped. Exine sculpture foveolate. (Plate 1)

Examined taxa: Acacia nilotica (L.) Delile 1b. Faidherbia subtype

Polyads slightly elliptic, large in size (41.8-51.8 Pm x 91.9-115.2 Pm), composed of 32 grains. 16 grains in the centre, placed in two planes, each plane eight grains surrounded by 16 peripheral grains. Pollen grains with indistinct aperture. Exine sculpture subrugulate. (Plate 2).

Examined taxa: Faidherbia albida (Delile) A. Chev. B. Single grains group 2. Triporate pollen type

Pollen grains small in size with mean polar axis 13.4 ±1.1 Pm and mean equatorial diameter 14.7 ±1.3 Pm, suboblate or oblate spheroidal in equatorial view, circular in polar view. Grains triporate, pores circular, sunken, annulate with microechinate operculum. Exine sculpture tectate, tectum densely spaced microechinate. This type characterized to Urticaceae (Urtica urnes L.), (Plate 3). 3. Tetra- Hexaporate pollen type

Pollen grains small in size with mean dimension 24.9 ±1.3Pm, spheroidal and circular in all views. Grains tetra to hexaporate, pores circular, raised, aspidate, with psilate and membranous-like operculum. Exine sculpture tectate, tectum verrucate with faint perforate. This type characterized to Fumariaceae (Fumaria bracteosa Pomel), (Plate 4) 4. Pantoporate pollen type

Pollen grains ranged in its size from small (13.4-25.1Pm in diameter) to medium (31.7-51.8Pm in diameter) or relatively large in size (75.2-85.2Pm in diameter), spheroidal and circular in all views, sometimes octagonal (in genus Alternanthera). Grains Pantoporate, pores ranged from 12 to many porate, circular in outline, flat or sunken, operculate or not operculate. Exine sculpture tectate to semitectate or intectate, tectum different patterns. This type represented by six pollen subtypes as follow:


Plate ( 1 ). Polyads pollen type ( Acacia pollen subtype)

Acacia nilotica (L.) Delile (Leguminosae)

1. SEM micrographs showing polyad in polar view, circular with 16 grains and Y-shaped colpi (scale bar = 10Pm)

2-3. SEM micrographs showing polyad in equatorial view, oblate (scale bar = 10Pm). 4. SEM micrographs showing foveolate sculpture (scale bar = 1Pm). 5-6. LM micrographs showing polyad in polar view (scale bar = 10Pm). 7-8. LM micrographs showing polyad in equatorial view (scale bar = 10Pm).


Plate ( 2 ). Polyads pollen type ( Faidherbia pollen subtype)

Faidherbia albida (Delile) A.Chev. (Leguminosae)

1-2. SEM micrographs showing polyad in polar view, ellipsoidal with 32 grains (scale bar = 10Pm)

3. SEM micrographs showing exine sculpture (scale bar = 10Pm). 4. SEM micrographs showing subrugulate sculpture (scale bar = 1Pm). 5-6. LM micrographs showing polyad in polar view (scale bar = 10Pm). 7. LM micrographs showing polyad in equatorial view (scale bar = 10Pm).


Plate ( 3 ). Triporate pollen type

Urtica urens L. (Urticaceae) 1-2. SEM micrographs showing pollen in equatorial view oblate-spheroidal or suboblate,

pore sunken, annulate with microechinate operculum, densely spaced microechinate sculpture (scale bar = 10Pm).

Plate ( 4 ): Tetra- Hexaporate pollen type

Fumaria bracteosa Pomel (Fumariaceae) 1-3. SEM micrographs showing pollen overall view spheroidal (scale bar = 10Pm) 4. SEM micrographs showing verrucate with faint perforate sculpture, pores raised,

aspidate and operculate (scale bar = 1Pm). 5-8. LM micrographs showing pollen overall view (scale bar = 10Pm).


4a. Persicaria pollen subtype: Pollen grains medium sized, mean dimension 45.8 ±3.8Pm, pores many

porate, sunken, filled the lumen of reticulum, not operculate. Exine sculpture semitectate, tectum lophoreticulate pattern, lophae with ±regular mural pattern, lumina polygonal with free standing columellae. This pollen subtype characterized to genus Persicaria (P. salicifolia (Brouss.ex Willd.) Assenov.), (Plate 5). 4b. Malvaceae pollen subtype

Pollen grains large in size, mean dimension 80.3 ±3.0Pm, pores many, flat, not operculate. Exine sculpture intectate, sharp echinate and interspinal area perforate, perforation densely spaced and uneven. This pollen subtype characterized to Malvaceae (Malva parviflora L.), (Plate 6). 4c. Silene-Vaccaria pollen subtype

Pollen grains medium sized, mean dimension 34.5 ±0.2 – 38.3 ±1.3Pm. Pores 12-many porate, ±sunken, operculate, operculum with 5-6 spinules. Exine sculpture tectate, tectum microechinate with sparsely spaced and even anulopunctate. This type characterized to genus Silene and genus Vaccaria (Silene rubella L. & Vaccaria hispanica (Mill.) Rauschert), (Plate 7). 4d. Thymelaeaceae pollen subtype

Pollen grains small sized, mean dimension 22.7 ±1.1Pm. Pores many porate, sunken and not operculate. Exine sculpture intectate, croton pattern. This pollen subtype characterized to Thymelaeaceae (Thymelaea hirsute (L.) Endl.), (Plate 8). 4e. Alternanthera pollen subtype

Pollen grains small sized, mean dimension 14.6 ±1.0Pm, octagonal in all views. Pores 12, each filled the lumen of reticulum, ± flat with striate membrane. Exine sculpture semitectate, metareticulate (fenestrate) pattern, reticulum with regular mural pattern, muri microechinate and faintly perforate. This pollen subtype characterized to genus Alternanthera ( A. sessilis (L.)DC.), (Plate 9). 4f. Amaranthus-Chenopodiaceae pollen subtype

Pollen grains small sized, mean dimension 18.6 ±1.1 – 24.5 ±1.3Pm. Pores many, sunken, operculate, operculum with many coarsely granules. Exine sculpture granulate – psilate to granulate – faint perforate. This pollen subtype characterized to genus Amaranthus and Family Chenopodiaceae (Amaranthus graecizans L., Arthrocnemum macrostachyum (Moric.)K. Koch, Beta vulgaris L., Chenopodium album L. & Suaeda aegyptiaca (Hasselq.) Zohary, (Plate 10)


Plate ( 5 ). Pantoporate pollen type (Persicaria pollen subtype)

Persicaria salicifolia (Brouss. ex Willd.) Assenov (Polygonaceae)

1-2. SEM micrographs showing pollen overall view spheroidal, many porate (scale bar = 10Pm)

3. SEM micrographs showing lophoreticulate pattern sculpture, pores sunken and filled the lumen of reticulum (scale bar = 1Pm).

4. SEM micrographs showing lumina polygonal with free standing collumellae (scale bar = 1Pm).

5-7. LM micrographs showing pollen overall view (scale bar = 10Pm).


Plate ( 6 ). Pantoporate pollen type (Malvaceae pollen subtype) Malva parviflora L. (Malvaceae)

1-2. SEM micrographs showing pollen overall view spheroidal, many porate (scale bar =

10Pm) 3. SEM micrographs showing sharp echinate-perforate sculpture (scale bar = 10Pm). 4. SEM micrographs showing perforation densely spaced and uneven, pores flat and not

operculate (scale bar = 1Pm). 5-8. LM micrographs showing pollen overall view (scale bar = 10Pm).


Plate ( 7 ). Pantoporate pollen type (Silene-Vaccaria pollen subtype) Silene rubella L. (Caryophyllaceae)

1-3. SEM micrographs showing pollen circular in all views with many porate,

pores sunken and operculate, operculum with 5-6 spinules (scale bar = 10Pm).

4. SEM micrographs showing microechinate with anulopunctate sculpture (scale bar = 1Pm) 5-8. LM micrographs, (overall view), scale bar = 10Pm


Plate ( 8 ). Pantoporate pollen type (Thymelaeaceae pollen subtype) Thymelaea hirsute (L.) Endl. (Thymelaeaceae)

1-3. SEM micrographs showing pollen overall view spheroidal, (scale bar = 10Pm) 4. SEM micrographs showing microechinate croton pattern sculpture (scale bar =

1Pm). 5-8. LM micrographs showing pollen overall view (scale bar = 10Pm).


Plate ( 9 ). Pantoporate pollen type (Alternanthera pollen subtype) Alternanthera sessilis (L.) DC. (Amaranthaceae)

1-3. SEM micrographs showing pollen octagonal in all views with 12 porate, pores

flat and striate membrane (scale bar = 10Pm). 4. SEM micrographs showing Metareticulate sculpture (scale bar = 1Pm). 5-8. LM micrographs, (overall view), scale bar = 10Pm


Plate ( 10 ). Pantoporate pollen type (Amaranthus-chenopodiaceae pollen subtype)

Chenopodium album L. (Chenopodiaceae)

1-3. SEM micrographs showing circular in all views with many porate, pores sunken and operculate, operculum with many granules (scale bar = 10Pm).

4. SEM micrographs showing granulate - psilate sculpture (scale bar = 1Pm) 5-8. LM micrographs, (overall view), scale bar = 10Pm

5. Pantobrevicolpate pollen type

Pollen grains relatively large in size, mean dimension 53.6 ±3.2Pm, spheroidal and circular in all views. Grains pantobrevicolpate, colpi short, fusiform. Exine sculpture tectate, tectum microechinate with sharp anulopunctate (perforation densely spaced and even). This type characterized to Portulacaceae (Portulaca oleracea L.), (Plate 11).


Plate ( 11 ). Pantobrevicolpate pollen type Portulaca oleracea (portulacaceae)

1-3. SEM micrographs showing pollen overall view spheroidal, colpi short and fusiform,

(scale bar = 10Pm) 4. SEM micrographs showing microechinate with sharp anulopunctate sculpture (scale bar

= 1Pm). 5-7. LM micrographs showing pollen overall view (scale bar = 10Pm).


6. Tricolpate pollen type Pollen grains small to medium sized with mean polar axis ranges from

14.2 ±0.9- 31.1 ±2.5Pm, mean equatorial diameter ranges from 14.6 ±0.9- 32.7 ±2.1Pm, spheroidal to oblate – spheroidal or suboblate in equatorial view and circular or obtusely triangular in polar view. Grains tricolpate, colpi apocolpate, fusiform to oblong, with acute to obtuse apex, colpi membrane scabrate to densely granules. Exine sculpture tectate to semitectate, tectum varies from reticulate to microreticulate-perforate or granulate-perforate. This type included four subtypes which can be distinguished as follows: 6a. Spergularia pollen subtype

Pollen grains small sized, mean polar axis 18.7 ±1.4Pm, mean equatorial diameter 19.9 ±1.6Pm, spheroidal or suboblate in equatorial view and circular in polar view. Colpi oblong with obtusely apex, colpi membrane densely granules. Exine sculpture tectate, tectum granulate -perforate. This pollen subtype characterized to genus Spergularia (S. marina (L.) Bessler), (Plate 12).

Plate ( 12 ). Tricolpate pollen type (Spergularia pollen subtype)

Spergularia marina (L.) Bessler (Caryophyllaceae)

1-2. SEM micrographs showing pollen in equatorial view, spheroidal or suboblate, colpi oblong with obtusely apex, (scale bar = 10Pm)

3. SEM micrographs showing pollen in polar view circular, (scale bar = 10Pm). 4. SEM micrographs showing granulate-perforate sculpture, (scale bar = 1Pm).


6b. Cruciferae pollen subtype Pollen grains small sized, mean polar axis ranges from 23.6 ±1.4- 23.9

±1.1Pm, mean equatorial diameter ranges from 23.7 ±1.2- 26.5 ±1.2Pm, spheroidal, oblate-spheroidal or suboblate in equatorial view and obtusely triangular in polar view. Colpi fusiform with acute apex, colpi membrane densely granules. Exine sculpture semitectate, tectum reticulate, reticulum with free standing columellae in the lumina. This pollen subtype characterized to Cruciferae (Brassica nigra (L.) Koch, Sinapis arvensis L.), (Plate 13).

Plate ( 13 ). Tricolpate pollen type (Cruciferae pollen subtype)

Brassica nigra (L.) Koch (Cruciferae) 1-2. SEM micrographs showing pollen in equatorial view, spheroidal or oblate spheroidal,

colpi fusiform with acute apex, (scale bar = 10Pm) 3. SEM micrographs showing pollen in polar view obtusely triangular, (scale bar = 10Pm). 4. SEM micrographs showing reticulate sculpture with free standing columellae in the

lumina, (scale bar = 1Pm). 5-7. LM micrographs showing pollen in equatorial view (scale bar = 10Pm). 8. LM micrographs showing pollen in polar view (scale bar = 10Pm).


6c. Tamaricaceae pollen subtype Pollen grains small sized, mean polar axis ranges from 14.2 ±0.9-16.9

±0.9Pm, mean equatorial diameter ranges from 14.6 ±0.9- 16.8 ±1.1Pm, spheroidal or subprolate in equatorial view and obtusely triangular in polar view. Colpi fusiform with acute apex, colpi membrane psilate-scabrate. Exine sculpture tectate, tectum microreticulate to perforate. This pollen subtype characterized to Tamaricaceae ( Tamarix nilotica (Ehrenb.)Bunge & T. tetragyna Ehrenb.), (Plate 14)

Plate ( 14 ). Tricolpate pollen type (Tamaricaceae pollen subtype)

Tamarix tetragyna Ehrenb. (Tamaricaceae)

1-2. SEM micrographs showing pollen in equatorial view spheroidal or subprolate, colpi fusiform with acute apex (scale bar = 10Pm)

3. SEM micrographs showing pollen in polar view obtusely triangular (scale bar = 10Pm).

4. SEM micrographs showing microreticulate to perforate sculpture (scale bar = 1Pm).


6d. Oxalidaceae pollen subtype Pollen grains medium sized mean polar axis 31.1 ±2.5 Pm, mean

equatorial diameter 32.7 ±2.1Pm, spheroidal in equatorial view and circular in polar view. Colpi fusiform with acute apex, colpi membrane densely granules to rugulate. Exine sculpture tectate, tectum microreticulate to perforate. This pollen subtype characterized to Oxalidaceae (Oxalis corniculata L.), (Plate 15).

Plate ( 15 ). Tricolpate pollen type (Oxalidaceae pollen subtype)

Oxalis corniculata L. (Oxalidaceae) 1-2. SEM micrographs showing pollen in equatorial view spheroidal, colpi fusiform with

acute apex (scale bar = 10Pm) 3. SEM micrographs showing pollen in polar view circular (scale bar = 10Pm). 4. SEM micrographs showing microreticulate sculpture, colpi membrane rugulate (scale bar

= 1Pm). 5-7. LM micrographs showing pollen in equatorial view (scale bar = 10Pm). 8. LM micrographs showing pollen in polar view (scale bar = 10Pm).


7. Tricolporate pollen type Pollen grains small to medium sized with mean polar axis ranges from

9.9 ±0.9- 37.6 ±2.5Pm, mean equatorial diameter ranges from 9.9 ±0.9- 27.9 ±2.1Pm, spheroidal to prolate – spheroidal or subprolate to prolate in equatorial view and circular or obtusely triangular in polar view. Grains tricolporate, colpori apocolpate sometimes syncolpate, very short or long, fusiform to oblong to slit-like, with acute to obtuse apex, colpi membrane psilate or granulate to coarsely rugulate, margo distinct or indistinct. Exine sculpture tectate or semitectate, tectum varies from perforate to reticulate or rugulate. This type included six subtypes which can be distinguished as follows: 7a. Melilotus-group pollen subtypes

Pollen grains small sized with mean polar axis ranges from 16.5 ±1.3- 24.2 ±1.0Pm, mean equatorial diameter ranges from 14.1 ±0.8- 24.7 ±1.4Pm, spheroidal to prolate – spheroidal or subprolate to prolate in equatorial view and subcircular in polar view. Colpori apocolpate, fusiform with acute apex, colpi membrane coarsely rugulate, margo distinct, psilate to perforate. Ora raised and lolongate. Exine sculpture tectate, tectum perforate to microreticulate. This subtype characterized to Melilotus (M. indicus (L.) All.), Alhagi (A. graecorum Boiss.) and Sesbania (S. sesban (L.) Merr.), (Plate 16) 7b. Vicia-group pollen subtype

Pollen grains medium sized with mean polar axis ranges from 26.3 ±0.9- 34.1 ±1.4Pm, mean equatorial diameter ranges from 20.6 ±1.0- 22.1 ±0.9Pm, prolate or subprolate in equatorial view and subcircular in polar view. Colpori apocolpate, oblong to slit-like with acute apex, colpi membrane indistinct, margo distinct, psilate to perforate. Ora slightly raised and lolongate. Exine sculpture tectate, tectum uneven reticulate-rugulate to reticulate-fossulate. This subtype characterized to Vicia (V. monantha Retz.) and Trifolium (T. resupinatum L.), (Plate 17). 7c. Lotus-group pollen subtype

Pollen grains small to medium sized with mean polar axis ranges from 16.1 ±1.0- 37.6 ±2.1Pm, mean equatorial diameter ranges from 11.1 ±0.7- 24.9 ±1.4Pm, spheroidal, prolate or subprolate in equatorial view and circular to obtusely triangular in polar view. Colpori apocolpate, oblong to slit-like with acute apex, colpi membrane psilate to granulate, margo indistinct. Ora not raised and lolongate. Exine sculpture tectate, tectum rugulate to rugulate-fossulate. This subtype characterized to Lotus (L.


pusillus Viv. & L. halophilus Boiss. & Spruner), Lathyrus (L. hirsutus L.) and Medicago (M. intertexta (L.) Mill var. ciliaris (L.) Heyn), (Plate 18).

Plate ( 16 ). Tricolporate pollen type (Melilotus-group pollen subtype)

Melilotus indicus (L.) All. (Leguminosae) 1-2. SEM micrographs showing pollen in equatorial view subprolate or prolate,

colpi narrowly fusiform with acute apex, margo distinct (scale bar = 10Pm) 3. SEM micrographs showing pollen in polar view subcircular, (scale bar = 10Pm). 4. SEM micrographs showing microreticulate sculpture and colpi membrane

rugulate, (scale bar = 1Pm). 5-7. LM micrographs showing pollen in equatorial view (scale bar = 10Pm). 8. LM micrographs showing pollen in polar view (scale bar = 10Pm).


Plate ( 17 ). Tricolporate pollen type (Vicia-group pollen subtype) Trifolium resupinatum L. (Leguminosae)

1-2. SEM micrographs showing pollen in equatorial view subprolate or prolate, colpi narrowly oblong with acute apex, margo distinct (scale bar = 10Pm)

3. SEM micrographs showing pollen in polar view subcircular, (scale bar = 10Pm). 4. SEM micrographs showing reticulate-fossulate sculpture, (scale bar = 1Pm). 5-7. LM micrographs showing pollen in equatorial view (scale bar = 10Pm). 8. LM micrographs showing pollen in polar view (scale bar = 10Pm).


7d. Salicaceae pollen subtype Pollen grains small sized with mean polar axis 20.8 ±1.6Pm, mean

equatorial diameter ranges from 17.1 ±1.4Pm, spheroidal, subprolate or prolate in equatorial view and obtusely triangular in polar view. Colpori syncolpate, fusiform with acute apex, colpi membrane granulate, margo distinct, psilate to perforate. Ora sunken and circular. Exine sculpture semitectate, tectum uneven reticulate with granules at lumina. This subtype characterized to Salicaceae (Salix mucronata Thunb.), (Plate 19). 7e. Polygonaceae-group pollen subtype

Pollen grains small to medium sized with mean polar axis ranges from 20.4 ±0.8- 30.0±1.2Pm, mean equatorial diameter ranges from 19.5 ±0.9- 26.4 ±1.2Pm, spheroidal to oblate – spheroidal or prolate in equatorial view and circular to subcircular in polar view. Colpori apocolpate, oblong to slit- like (very short and fusiform in genus Emex) with acute apex, colpi membrane scabrate sometimes indistinct, margo indistinct. Ora sunken and lolongate. Exine sculpture tectate, tectum fossulate-perforate to microperforate with densely or sparsely spaced granules. This subtype characterized to Aizoaceae (Mesembryanthemum crystallium L.) and Polygonaceae except Persicaria (Emex spinosa (L.) Campd, Polygonum equsetiforume Sm., Rumex dentatus L. & R. pictus Forssk.), (Plate 20 & 21). 7f. Zygophyllaceae-group pollen subtype

Pollen grains small sized with mean polar axis ranges from 9.9 ±0.3- 18.8 ±1.4Pm, mean equatorial diameter ranges from 9.9 ±0.9- 16.3 ±1.3Pm, spheroidal, subprolate or prolate in equatorial view and obtusely triangular or circular in polar view. Colpori apocolpate, widely oblong with obtuse apex or fusiform with acute apex, colpi membrane psilate, margo distinct, psilate to perforate. Ora not raised and lolongate or lalongate. Exine sculpture tectate, tectum perforate to microreticulate. This subtype characterized to Capparaceae (Capparis decidus (Forssk.) Fdgew.), Salvadoraceae (Salvadora persica L.) and Zygophyllaceae (Fagonia arabica L. & Zygophyllum coccineum L.), (Plate 22).


Plate ( 18 ). Tricolporate pollen type (Lotus-group pollen subtype) Lotus halophilus Boiss. & Spruner ( Leguminosae)

1-2. SEM micrographs showing pollen in equatorial view, prolate, colpi slit-like

with acute apex, (scale bar = 10Pm) 3. SEM micrographs showing pollen in polar view circular, (scale bar = 10Pm). 4. SEM micrographs showing rugulate-fossulate sculpture, (scale bar = 1Pm). 5-7. LM micrographs showing pollen in equatorial view (scale bar = 10Pm). 8. LM micrographs showing pollen in polar view (scale bar = 10Pm).


Plate ( 19 ). Tricolporate pollen type (Salicaceae pollen subtype) Salix mucronata Thunb. (Salicaceae)

1-2. SEM micrographs showing pollen in equatorial view subprolate or prolate, colpi

fusiform with acute apex, margo distinct (scale bar = 10Pm) 3. SEM micrographs showing pollen in polar view obtusely triangular, syncolpate (scale

bar = 10Pm). 4. SEM micrographs showing uneven reticulate with granules at lumina sculpture (scale bar

= 1Pm). 5-7. LM micrographs showing pollen in equatorial view (scale bar = 10Pm). 8. LM micrographs showing pollen in polar view (scale bar = 10Pm).


Plate (20). Tricolporate pollen type (Polygonaceae-group pollen subtype)

Polygonum equsetiforme Sm. (Polygonaceae) 1-2. SEM micrographs showing pollen in equatorial view subprolate or prolate,

colpi oblong with acute apex, margo indistinct (scale bar = 10Pm) 3. SEM micrographs showing pollen in polar view subcircular, (scale bar = 10Pm). 4. SEM micrographs showing microperforate-granulate sculpture (scale bar =

1Pm).

Rumex pictus Forssk. (Polygonaceae)

1-2. SEM micrographs showing pollen in equatorial view suboblate or oblate spheroidal, slit-like colpus and fossulate-perforate with granules sculpture (scale bar = 10Pm)


Plate (21). Tricolporate pollen type (Polygonaceae-group pollen subtype)

Emex spinosa (L.) Campd (Polygonaceae) 1-2. SEM micrographs showing pollen in equatorial view suboblate or spheroidal, colpi

short and fusiform with acute apex, margo indistinct (scale bar = 10Pm) 3. SEM micrographs showing pollen in polar view circular, (scale bar = 10Pm). 4. SEM micrographs showing fossulate-perforate with densely granules sculpture (scale bar

= 1Pm).


Plate ( 22 ). Tricolporate pollen type (Zygophyllaceae-group pollen subtype)

Salvadora pesica Wall. (Salvadoraceae) 1-2. SEM micrographs showing pollen in equatorial view spheroidal or subprolate, colpi

widely oblong with obtuse apex, margo distinct and perforate (scale bar = 10Pm) 3. SEM micrographs showing pollen in polar view obtusely triangular (scale bar = 10Pm). 4. SEM micrographs showing perforate sculpture (scale bar = 1Pm).

Zygophyllum coccineum L. (Zygophyllaceae)

1-2. SEM micrographs showing pollen in equatorial view spheroidal or subprolate, colpi fusiform with acute apex, margo distinct and psilate. Sculpture microreticulate (scale bar = 10Pm)


Discussion Scanning electron micrographs greatly enhanced the characterization

and interpretation of pollen morphological features (Sowunmi, 1995). The SEM of the 41 taxa from different families of subclass Archiclamydeae in Faiyum province provided further information on the pollen morphology and complemented the light microscopic observations. Number and shape of apertures, size and shape of pollen grains and exine patterns were of great diagnostic significance among the examined taxa.

On the basis of pollen units, two main groups are recognized viz. complex group and single group. The complex group is represented by a unique major pollen type known as polyads type in which the pollen grains released in groups 16-32 grains, this type restricted to subfamily Mimosoideae and is further subdivided according to the number of grains into two subtypes viz. Acacia pollen subtype (16 grains in each polyad) and Faidherbia pollen subtype (32 grains in each polyad).

According to the number and shape of apertures, the single group (monads) comprises six pollen types of which triporate, tetra-hexaporate and pantobrevicolpate types are characterized to families Urticaceae, Fumariaceae and Portulacaceae respectively.

The pollen grains of pantoporate type are characterized by having spheroidal shape with different size and exine sculpture patterns. Six families are included in this type distributed in six subtypes viz. Malvaceae, Thymelaeaceae, Chenopodiaceae, Amaranthaceae, Caryophyllaceae and Polygonaceae (Persicaria).

Among the under investigation, Pollen of Malvaceae is the largest one, its dimension ranges from 75.2-85.2Pm, with sharp echinate tectum (Malvaceae pollen subtype) whereas the pollen of Thymelaeaceae is characterized by croton pattern tectum (Thymelaeaceae pollen subtype).

Amaranthaceae represented here by two genera viz. Amaranthus (subfamily Amaranthoideae) and Alternanthera (subfamily Gomphrenoideae). Both genera produce two different subtypes of pollen grains, the Alternanthera pollen subtype characterized by octagonal pollen grains in all views, pores 12 and exine sculpture metareticulate pattern; while the Amaranthus pollen subtype has circular pollen grains in all views, pores more than 12 and exine sculpture granulate. These results agree with that of Erdtman 1952, Nowick & Skvarla 1979 and Perveen & Qaiser 2002.

Within the order Caryophyllales, Chenopodiaceae regarded by Bittrich 1993 as a sister group with Amaranthaceae, both form a monophyletic


lineage (Downie & Palmer 1994, Manharht & Rettig 1994, Rodman 1994). Hence, the ultrastructure of pollen grains of the studied genera of Chenopodiaceae, shows uniform type characteristics and resembles Amaranthaceae (Amaranthus pollen subtype) in several aspects such as the aperature structure, exine sculpture and pollen shape and size. This was also noted by (Tsukada 1967, Youngjae & Lee 1995 and Toderich et al. 2010).

Pollen grains of Persicaria (family Polygonaceae) characterized by the lophoreticulate sculpture whereas the remaining examined genera of family Polygonaceae characterized by tricolporate apertures and perforate-scabrate sculpture (tricolporate pollen type) sharing family Aizoaceae in the same pollen subtype. As well as Caryophyllaceae is a europalynous family, both Silene and Vaccaria belonging to pantoporate pollen type with microechinate-anulopunctate sculpture (Silene-Vaccaria pollen subtype). While Spergularia belong to tricolpate pollen type with granulate-perforate sculpture (Spergularia pollen subtype). Nowicke & Skvarla 1977 reported that Polygonaeae is a large and palynological diverse family but the palynological study revealed no pollen types similar to these in the Caryophyllaceae.

Tricolpate pollen type represented here by four pollen subtypes of which pollen of Cruciferae is characterized by reticulate sculpture, whereas both Tamaricaceae and Oxalidaceae have the same exine sculpture (microreticulate to perforate) but differ in pollen size, shape and sculpture of colpi membrane.

The pollen of tricolporate type is quite heterogeneous, most of the examined taxa of this type belong to the subfamily Papilinoideae, and these taxa show a wide range of variation in pollen characteristics. On the basis of pollen shape and size, aperture characters and exine sculpture, six pollen subtypes are recognized, of which three subtypes included eight genera of papilionoideae viz. Melilotus group pollen subtype, Vicia group pollen subtype and Lotus group pollen subtype. Pollen of Melilotus group is easily distinguished by its fusiform colpori, coarsely regulate colpi membrane and tectum perforate to microreticulate. This subtype comprises three genera (Sesbania, Alhagi and Melilotus) belonging to three tribes: Robinieae, Galegeae and Trifolieae respectively. Vicia group pollen subtype is characterized by distinct margo, psilate to perforate and uneven reticulate- rugulate to reticulate-fossulate tectum. This pollen subtype comprises two genera (Vicia and Trifolium) belonging to two tribes: Vicieae and Trifolieae respectively. While the pollen of Lotus group is characterized by indistinct


margo and regulate-fossulate tectum. This pollen subtype is found in three genera (Lotus, Lathyrus and Medicago) distributed in three tribes: Loteae, Vicieae and Trifolieae respectively. Striking variations were found in pollen characters within this subfamily indicated that it is a eurypalynous; and hence palynology is significant within this subfamily at the tribal and subtribal levels, but less marked at the genera levels. These results also cited by Erdtman 1952, Ferguson & Skvarla 1981 and Perveen & Qaiser 1998.

The remaining three subtypes represented the pollen of six families viz. Salicaceae pollen subtype, Polygonaceae group pollen subtype and Zygophyllaceae group pollen subtype. The former subtype distinguished by reticulate tectum, with free standing columellae in lumina, pollen small sized and colpus syncolpate.

Although the other two pollen subtypes are fairly uniform in their pollen characteristics. On the basis of exine sculpture and colpori characters, the pollen of Polygonaceae group can be distinguished by tectum fossulate- perforate to microperforate with densely or sparsely spaced granules and colpi slit-like with indistinct margo. While, pollen of Zygophyllaceae group is characterized by perforate to microreticulate tectum and colpi widely oblong or fusiform with distinct margo.

In conclusion, SEM based pollen characters are found to be useful in identification and discrimination of taxonomically related taxa. The taxa investigated here illustrate a highly pollen morphological variations especially among the taxa related to families: Leguminosae, Caryophyllaceae, Amaranthaceae and Polygonaceae. The key to identify pollen types and examined taxa given in the present investigation can be additional tool for taxonomic identification. In the future, an investigation of the taxa belong to the other subclass (Sympetalae) will be taken into consideration by the authors. Key based on the morphological characters of pollen grains of investigated taxa: 1- Pollen in polyads………………………………………………..………. 2

-Pollen in Monads…………………………………...…………...……… 3 2- Polyads medium sized (23.4-26.7Pm x 36.7-43.4Pm), composed of 16

grains, pollen grains colporate, colpi Y-shaped, exine sculpture foveolate ……………………………………………………..…. Acacia

- Polyads large in size (41.8-51.8 Pm x 91.9-115.2Pm), compoded of 32 grains, pollen grains indistinct aperture, exine sculpture subrugulate ………………………….………………………………….. Faidherbia


3- Pollen aperture is porate ………………………………………….…… 4 - Pollen aperture is colpate or colporate ………………………….11

4- Pollen aperture tri- or tetra-hexaporate ……………………………. 5 - Pollen aperture pantoporate …………………………………………... 6

5- Pollen aperture triporate, pores circular with microechinate annulus, exine sculpture densely spaced micrechinate …………...….. Urticaceae

- Pollen aperture tetra to hexaporate, pores circular with psilate aspides, exine sculpture verrucate with faint perforate ……..…..... Papaveraceae

6- Pollen dimension ranges from 31.7-85.2Pm ……………………. 7 - Pollen dimension ranges from 13.4-30.1Pm ………………………….. 9

7- Exine sculpture lophoreticulate with free standing columellae in lumina …………………………………………………….……… Persicaria

- Exine sculpture microechinate or sharp echinate with perforate or anulopunctate ………………………………………………………. 8

8- Pollen dimension ranges from 75.2-85.2Pm, exine sculpture sharp echinate, perforation densely spaced and uneven ………....... Malvaceae

- Pollen dimension ranges from 31.7-40.9Pm, exine sculpture microechinate, perforation sparsely spaced and circular (anulopunctate) …………………….……….………………… Silene-Vaccaria group 9- Exine sculpture is croton pattern ………………….…… Thymelaeaceae

- Exine sculpture is metareticulate (fenestrate) pattern or granulate .......................................................................................................... 10

10- Exine sculpture metareticulate (fenestrate) pattern, with microechinate and perforate muri. Pores 12, striate membrane. Pollen octagonal in all views ………………………………...……………….. Alternanthera

- Exine sculpture granulate pattern. Pores more than 12 with coarsely granules operculum. Pollen circular in all views …………………….……………… Amaranthus & Chenopodiaceae

11- Pollen aperture tri- or panto-colpate ……………………………. ...... 12 - Pollen aperture tri-colporate ……………………………………. 16

12- Pollen aperture pantobrevicolpate ………………….. Portulacaceaea - Pollen aperture tricolpate ………………………………………. 13

13- Exine sculpture granulates and perforate ............................ Spergularia - Exine sculpture reticulate or microreticulate ……………………. 14

14- Exine sculpture reticulate, reticulum with free standing columellae in lumina ……………………………….…………………… Cruciferae

- Exine sculpture microreticulate to perforate……………………. 15


15- Pollen grains small sized (13.4-18.4Pm x 15-19.2Pm), spheroidal or subprolate. Colpi membrane psilate to scabrate ………... Tamaricaceae

- Pollen grains medium sized (its dimension 23.4-36.7Pm), spheroidal. Colpi membrane densely granulate .............................…… Oxalidaceae 16- Exine sculpture rugulate to rugulate-fossulate ..………...... Lotus group

- Exine sculpture otherwise …………………………………………… 17 17- Exine sculpture microreticulate to fossulate-perforate or microperforate.

Supratectum psilate or best with granules …………………………… 18 - Exine sculpture reticulate ……………………………………….21

18- Supratectum best with granules ………….. Aizoaceae & Polygonaceae - Supratectum psilate …………………………………………………. 19

19- Colpi membrane coarsely regulate ……...……….…… Melilotus group - Colpi membrane psilate …………………………………………. ....... 20

20- Colpus fusiform with acute apex …... Capparaceae & Zygophyllaceae - Colpus widely oblong with obtuse apex ……………… Salvadoraceae

21- Exine sculpture reticulate with free standing columellae in lumina. Grains small sized (16.7-23.4Pm x 15.0-20.0Pm). Colpus fusiform, syncolpate with perforated margin ……….……………… Salicaceae

- Exine sculpture uneven reticulate-rugulate to reticulate-fossulate. Grains medium sized (25.1-36.7Pm x 19.2-23.4Pm). Colpus oblong, apocolpate with thick & psilate to perforate margin …………… Trifolium & Vicia

Acknowledgement.

Grateful thanks to Dr. Amal Hossny (Botany Department, Faculty of Science, Cairo University) for valuable comments and critical revision of the manuscript.

References

Abd el Ghani, M. (1985). Comparative study on the vegetation of Bahariya

and Farafra Oases & Fayoum region. Ph D. thesis, Fac. of Sci. Univ. of Cairo.

Ayyad, S.M., Moore, P.D. and Zahran, M.A. (1992b).Modern Pollen rain studies of the Nile Delta, Egypt. New Phyotolgist, 2: 663-675.

Bittrich, V. (1993). Introduction to Centrospermae.- In: The Families and Genera of vascular plants (ed. K. Kubitzki, pp. 13-19.) – Springer, Berlin.


Boulos, L. (2009). Flora of Egypt Checklist, revised annotated edition, Al Hadara publishing. Egypt.

Downi, S.R. & Palmer, J.D. (1994). Phylogenetic relationships using restriction site variation of the chloroplast DNA inverted repeat.- In: Caryophyllales. Evolution & Systematics (ed. H.-D. Behnke & T.J. Mabry), Pp. 223-233.- Springer, Berlin.

El Fayoumi, H. (1996). Plants life in Fayoum area, Reconstruction study of the Natural vegetation using pollen analytical methods. Ph D. thesis, Fac. of Sci. Univ. of Cairo.

El Hadidi, M.N. (2000). The main features of the Natural Vegetation. In M.N. El Hadidi (ed.).- Flora Aegyptiaca 1(1)- The Palm Press, Cairo, Egypt.

-------------- & El Fayoumi, H.H. (1997). Palynological studies in El Gharag El Sultani area of the Faiyum depression, Egypt. 1- Pollen classes and Pollen types. Taeckholmia 17: 61-90.

El Shenbary, S.H. (1985). A study of recent changes in vegetation composition in the north-western coastal desert of Egypt in Burg El- Arab area. MSc thesis, Tanta University, Egypt.

Engler, A. (1964). Syllabus der pflanzenfamilien. (ed. 12) by H. Melchior Berlin.

Erdtman, G. (1952). Pollen Morphology and Plant Taxonomy. Angiosperms. Chronica Botanica Co., Waltham, Massachusettes.

------------ (1960). The acetolysis method.- Svensk Botanisk Tidskrift. 54: 561-564.

Ferguson, I.K. & Skvarla, J.J. (1981). The pollen morphology of the subfamily Papilionoideae (Leguminosae). In R.M. Polhill & P.H. Raven (eds.) Advances in legume systematics. Pp. 859-896. Royal Botanic Gardens, Kew, England.

Manhart, J.R. & Rettig, J.H. (1994). Gene sequence data.- In: Caryophyllales. Evolution and Systematics (ed. H.-D. Behnke & T.J. Mabry), Pp. 235-246.- Springer, Berlin.

Mehringer, J.R., Petersen, K.L. & Hassan, F.A. (1979). A pollen record from Birket Qarun and the recent history of the Fayoum, Egypt. Quaternary Research 2: 238-265.

Nowicke, J.W. & Skvarala, J.J. (1977). Pollen morphology and the Relationship of the Plumbaginaceae, Polygonaceae and the Primulaceae to the order Centrospermae. Smithsonian Contrib. Bot., 37: 1- 64.


----------------------------------------- (1979). Pollen morphology: The potential influence in higher order systemstics. Ann. Missouri Bot. Gard., 66: 633-699.

Perveen, A. & Qaiser, M. (1998). Pollen Flora of Pakistan. VIII. Legumin- osae (subfamily: Papilionoideae). Tr. J. of Botany., 22: 73-91.

---------------------------- (2002). Pollen Flora of Pakistan. XVIII. Amaranthaceae. Pak. J. Bot., 34(4): 375-383.

Praglowski, J. & Punt, W. (1973). An elucidation of the microreticulate structure of the exine. Grana, 13: 45-50

--------------- & Raj, B. (1979). On some pollen morphological concepts. Grana, 18: 109-113.

Punt, W., Hoen, P.P., Blackmore, S., Nilsson, S. & Le Thomas, A. (2007). Glossary of pollen and spores terminology. Rev. palaeobot. Palynol., 143(1-2): 1-81.

Ritchie, J.C. (1985). Modern pollen spectra from Dakhla Oasis, Western Egyptian desert. Grana, Uppsala, 1984, Pp. 1-6.

Rodman, J.E. (1994). Cladistic and Phenetic studies.- In: Caryophyllales. Evolution & Systematics (ed. H.-D. Behnke & T.J. Mabry), Pp. 279- 301.- Springer, Berlin.

Saad, S.I. & Sami, S. (1967). Studies of pollen and spores content of the Nile Delta deposits (Berenbal Region). Pollen et spores, 9: 467-503.

Sowunmi, M.A. (1995). Pollen of Nigerian. II. Woody species.- Grana, 34: 120-141.

Täckholm, V. (1974). Students' Flora of Egypt, ed.2. - Cairo University. Toderich, K.N., Shuyskaya E.V., Ozturk, M., Juylova, A. & Gismatulina, L.

(2010). Pollen morphology of some Asiatic species of Genus Salsola (Chenopodiaceae) and its taxonomic relationships. Pak. J. Bot., special issue, 42: 155-174.

Tsukada, M. (1967). Chenopod and Amaranth pollen. Electron Microscopic Identification. Scien. USA, 17: 80-82.

Youngjae, C. & Lee, S. (1995). Pollen morphology of some Korean Chenopodiaceae. Korean J. Plant Taxon., 25(4): 255-276.

Zahran, M.A. & Willis, A.J. (1992). The Vegetation of Egypt. 1st ed. Chapman & Hall, London.

--------------------------------- (2009). The Vegetation of Egypt. 2nd ed. Springer Science.

the pollen flora of faiyum, egypt i- archichlamydeae · 2020. 9. 3. · 2 eman m. shamso and fujiki...

Documents